USZO403 COURSE-10

Comparative Embryology, Aspects of Human Reproduction Pollution and Its Effect

on Organisms
Unit 1: Comparative embryology

Objective: To acquaint the learner with key concepts of embryology.

Desired Outcome: Learner will be able to understand and compare the different pre-embryonic stages

Learner will be able to appreciate the functional aspects of extra embryonic membranes
and classify the different types of placentae.

Introduction

Like growth, metabolism and irritability, reproduction is one of the properties of the living organisms.
Reproduction is the process of multiplication of individuals in the population belonging to a species. It
ensures the form and function of an organism. Animals have adapted for the multiplication and their
survival with the process of reproduction. Reproduction is basically classified as asexual and sexual types.
The sexual reproduction involves gametogenesis (formation of sex cells or gametes like sperms and ova
in germinal epithelium of testis and ovary respectively), fusion of gametes (fertilization), egg formation
and the embryonic development (zygote transformations). Development of embryo plays very important
role in reproduction. Also it is known that the embryonic development play important role to know the
evolutionary tract of the species (ontogeny repeats the phylogeny). Based on the embryonic evidences
animals are classified into protostomes and deuterostomes. Protostomes are inclusive of invertebrates
which develop mouth from blastopore (poriferans to molluscans) whilst the deuterostomes develop mouth
opposite to blastopore by forming separate opening and blastopore develops into anus (echinoderms
onwards). The higher vertebrates are further classified into anamniotes and amniotes as they do not or do
have the extraembryonic membrane known as amnion respectively.

1.1. Types of Eggs: Based on Amount and Distribution of Yolk

The primitive aquatic forms have adapted to reproduce in aquatic medium which is best suited for their
embryonic developments in due course of evolution. With the evolution the higher forms, like amniotes
(reptiles, birds and mammals), have adapted to live in dry terrestrial environment. However, in terrestrial
animals, who have internal fertilization but the external development of embryo (outside the body of
mother), their ova get transformed into shelled eggs which carry huge amount of yolk. In placental animals
the yolk is present in negligible amount since the embryo is nurtured through feto-maternal association of
tissue called placenta. Although yolk is an essential food reservoir playing important role in development
of the embryo, it has been observed that it causes the polarity in the egg and hinders the development of
embryo. The area of heavy deposition of yolk is therefore known as vegetal pole and on opposite side
with nucleus and concentrated cytoplasm is known as animal pole (Fig. 1.1). The animal pole inside the
egg is vital end where metabolism takes place for the embryonic development.
 Fig. 1.1: A schematic diagram showing animal and vegetal poles of polarized egg of frog.

Dependent on the adaptive radiation, animals have developed various methods of reproduction. The
development of ova or egg is not an exception. The ova and eggs, by and large, are commonly referred as
‘eggs’ and are classified into categories further based on the amount and allocation of yolk within the egg.

Fig. 1.1.a.: Classification of eggs on the bases of amount and distribution of yolk.
1.1.1. Classification of eggs based on amount of yolk

Based on the amount of yolk present, the eggs are classified into two major groups as oligolecithal and
polylecithal eggs which are further subdivided into alecithal, microlecithal, mesolecithal and
macrolecithal (megalecithal) respectively.

A. Oligolecithal Eggs:

Oligolecithal eggs contain negligible amount of yolk and hence they are not capable of complete
development of embryo and are subjected to undergo several metamorphic stages like larvae, pupae and
instars (invertebrates) or they acquire the support of mother by forming placenta (mammals) for further
development. These types of eggs undergo holoblastic cleavage since there is no hindrance due to yolk.

i. Alecithal Egg

Alecithal egg is characterized by its negligible amount of yolk. In such a case the embryo needs to be
implanted immediately after fertilization to form placenta. Placenta is a union of maternal and foetal
tissues formed inside the uterus of mother for the exchange of gases and nutrients so as to execute complete
development of the embryo e.g. Eutherians (placental mammals). Eutherians therefore show direct
development.

ii. Microlecithal Egg

This type of egg contains very little and mostly evenly distributed yolk. e.g. insects, Amphioxus and
tunicates. Therefore in insects the development is indirect and embryo undergoes stages of development
like larva which is allowed to feed extensively to acquire the energy essential for its further development
known as metamorphosis. Tunicates also form a short living tadpole larva before developing into an
adult. While in Amphioxus the development undergoes rapidly to cope with the meager yolk reserve in
the egg.

B. Polylecithal Eggs:

Generally oviparous animals produce polylecithal eggs. These eggs are heavily laden with yolk enough
for the complete development of the embryo before hatching. They are further classified as mesolecithal
and macrolecithal types.

i. Mesolecithal Egg

This type of egg contains moderate amount of yolk which is largely confined to vegetal pole. Animal pole
has cytoplasm and nucleus. The mesolecithal eggs undergo holoblastic cleavage but comparatively
slower, particularly in vegetal pole, than that of microlecithal eggs e.g. Petromyzon, Dipnoi (lungfish) and
amphibians.
ii. Macrolecithal Egg

Macrolecithal or megalecithal egg contains large amount of yolk. Terrestrial animals like reptiles, birds
and some mammals lay eggs with huge amount of yolk. This ensures a complete growth of embryo inside
the egg itself before it hatches out. Due to the large amount of yolk the cleavage is of meroblastic type
e.g. bony fishes, amphibians, reptiles and birds and monotremes (egg laying mammals).

1.1.2. Classification of eggs based on distribution of yolk

As the eggs of animals store different amount of yolk, there is a variation in its distribution in the egg
proper. Microlecithal eggs show even distribution of the yolk while in the macrolecithal eggs it may be
pushed to one end. Depending upon the distribution of yolk, the eggs are classified as isolecithal,
telolecithal and centrolecithal.

A. Isolecithal or homolecithal Egg

Isolecithal eggs contain very little amount of yolk which is distributed uniformly in the cytoplasm. These
eggs undergo holoblastic cleavage e.g. echinoderms, Amphioxus, mammals etc.

B. Telolecithal Egg

The eccentric distribution of yolk defines them as telolecithal eggs. These eggs contain very high amount
of yolk usually concentrated at the vegetal pole. Therefore the telolecithal eggs show pronounced polarity.
The vegetal pole is large and prominent in telolecithal eggs; hence cleavage is of meroblastic type (partial).
These eggs are further sub-divided as slightly telolecithal, moderately telolecithal or pronounced
telolecithal type.

a. Slightly telolecithal egg: It has small quantity of yolk which is distributed unevenly. The vegetal
hemisphere has the highest concentration and the animal hemisphere the lowest e.g. eggs of fishes. In
this case nucleus is placed in the centre. e.g. eggs of fishes.
b. Moderately telolecithal egg: It contains a moderate quantity of yolk which is distributed unevenly.
Due to high concentration of yolk in the vegetal hemisphere, the nucleus is shifted more towards the
animal hemisphere e.g. egg of amphibian.
c. Pronounced telolecithal egg: In this type of egg, due to the heavy deposition of yolk, the entire
vegetal hemisphere and a major portion of the animal hemisphere is occupied by yolk. Due to this
extremely uneven distribution of yolk, the ooplasm and nucleus are displaced towards the animal pole
e.g. egg of reptiles and birds.
C. Centrolecithal Egg

The centre of the egg is occupied by nucleus and a small amount of cytoplasm which is covered by a thick
layer of yolk. At the periphery the yolk is surrounded by a thin layer of cytoplasm which is connected by fine
strands of cytoplasm to central vital part e.g. arthropods and some coelenterates.

Fig. 1.1.b.: Diagrammatic representation of classification of eggs on the bases of amount and
distribution of yolk.

1.1.3. Mosaic and regulative eggs:

A. Mosaic or Determinate Egg: Every part to be developed of the future embryo is predetermined
in egg itself before or at the time of fertilization. If any portion is removed from the egg then the
related organ does not develop in that embryo. Such an egg is called mosaic or determinate egg
e.g. Polychaete, Nimertine, Annelids, Molluscs and Ascidians.

B. Regulative or Indeterminate Egg: The eggs in which developmental potentialities for different
organs of future embryo are not predetermined, are referred to as regulative or indeterminate eggs.
In regulative eggs the determination occurs only after third cleavage. If the blastomere of such an
egg is removed before the third cleavage, it develops into a complete embryo. In humans identical
twins develop due to such an attribute of the egg.

Eggs are either shelled or non-shelled. The animals which lay eggs on dry land, develop calcareous shell
are known as cleidoic egg e.g. reptiles and birds and the animals which exhibit internal development of
the embryo, lay their eggs without the shell, such eggs are referred as non-cleidoic eggs e.g. mammals.

1.2. Structure and Types of Sperms

Sperms or spermatozoa (singular: spermatozoon) are produced by the germinal epithelium of seminiferous
tubules by the process of spermatogenesis. Sperm is a means of transport of genetic material from male
to the oocyte of a female. It carries haploid set of chromosomes. It is short lived active gamete which has
to find the egg for fusion within a minute most of the time or else they perish.
 Fig.
The mature sperm cell is slender; in the
middle part, the mitochondria are thick
and ring-shaped. The DNA in the nucleus
is maximally condensed.

A. Head
B. Neck
C. Midpiece
D. Principal piece
E. End piece

Details of organelles of sperm

1. Plasma membrane
2. Outer acrosomal membrane
3. Acrosome
4. Nucleus
5. Proximal centriole
6. Rest of the distal centriole
7. Helix of Mitochondria

The sperm is microscopic structure and

measures about 65 µm in length in
humans (head 5 µm, neck=1 µm;
midpiece= 5 µm and tail= 55 µm).

Fig. 1.2.: Structure of sperm

1.2.1. Basic structure of sperm:

Although the structure of sperm varies from species to species, it is significantly similar in vertebrates.
The structure of a typical mammalian sperm is basically differentiated into head, neck and tail. Tail is
further differentiated into mid-piece, principle piece and end piece.

Head of spermatozoon

Head is the anterior most part of sperm and is made of a cap-like acrosome and nucleus.

Acrosome

The anterior end of sperm head is a crescentic, cap-like structure called acrosome. The shape and size of
the acrosome vary among different species. The acrosome enveloped by an acrosomal membrane and it
contains certain acrosomal polysaccharides viz. galactose, mannose, fructose, and hexosamine and many
enzymes, especially hydrolases. It also contains two proteolytic enzymes (called as hyaluronidase and
acrosin) which help in the entry of sperm into the ovum during fertilization.
Nucleus

The nucleus of the sperm occupies major part of the head. It comprises of DNA and nuclear proteins and
thus is responsible for the transmission of hereditary characters.

Neck

Neck is a constricted junction between the head and mid-piece. It carries anterior proximal centriole and
the posterior distal centriole. The distal centriole gives rise to central axial filaments called axoneme made
of microtubules to support the tail region which is used for locomotion. Neck also contains two or three
mitochondria and which lie in continuation with the anterior mitochondria of mid-piece.

Tail of spermatozoon

Tail or flagellum is the locomotory organelle of sperm and its length is approximately three fourth the
total length of the sperm. It has three parts viz. mid-piece, principal piece and end piece. The sperm moves
in a fluid medium inside the female genital tract by the undulating movement of the tail.

Mid-piece

Mid-piece lies between neck and principal piece of tail. It is made of an axial filament surrounded by helix
of mitochondria and little amount of cytoplasm. Mitochondria in mid-piece play an important role of
power house* to enable the tail (flagellum) to perform locomotion so as to reach the egg.

Principle piece

It is the second part of the tail consisting of the axoneme having microtubules arranged as 9 doublets
peripherally and 2 single ones in the center (typical 9+2 arrangement of an axoneme). Surrounding this
arrangement of microtubules is a sheath made of nine ring fibres. The fibrous sheath terminates at the
commencement of the end piece.

End-piece

The end piece has axoneme extending in it but is without fibrous sheath. The number of microtubules is
reduced merely to a pair of singlet microtubules at its terminal end.

1.2.2.Types of sperms in animals:

The structure of sperms varies in both invertebrates and vertebrates. The size of sperm ranges from 0.018
mm in Amphioxus to as large as 2.25 mm in toad. The shape of the head also varies from species to species.
It is branched or star-shaped in crustaceans and salamander; spherical in teleosts; lance-shaped in
amphibians and domestic fowl; hook-shaped in mouse and rat; spoon-shaped in several mammals
including man.

*
Mitochondria generate energy through oxidative phosphorylation of ATP.
Furthermore, the sperms may be flagellate or non-flagellate (Ascaris and some crustaceans). Flagellate
sperms may be uniflagellate having a single flagellum (many animals), biflagellate having two flagella
(toad fish and flatworm) and multiflagellate having many flagellae (some crustaceans and cladocerans).
In uniflagellate sperms the head differ in shape according to species e.g. rat has sickle shaped, amphioxus,
bull, rabbit, horse and humans have club shaped whereas domestic fowl has lancet shaped head of the
sperm.

Types of sperms in crustaceans

Fig. 1.2.2. Types of sperms in animals

1.3. Types of Cleavage- Holoblastic and Meroblastic

Cleavage:

Fertilized egg is a single-cell structure which gives rise to a multi-cellular embryo through the process of
cleavage. During cleavage, the zygote divides repeatedly by mitosis into progressively smaller cells called
blastomeres. Cleavage is characterized by rapid and successive mitotic divisions without any intervening
periods of growth. Therefore, blastomeres do not increase in size and the resulting blastomeres are only
half the original size. Thus, cleavage begins with one very large cell and ends with a large number of
smaller cells. Furthermore, since there is no growth in the size of the blastomeres, the total size and volume
of the embryo remains the same. The ratio of nucleus to cytoplasm is very low at the beginning of cleavage,
but at the end of cleavage (blastula stage); it is brought to the level found in ordinary somatic cells. Early
cleavage divisions occur synchronously i.e. all blastomeres present divide simultaneously and result in
the formation of a solid ball of cells called morula. But such synchrony usually disappears after a few
dozen blastomeres have been formed and a cavity appears which is called blastocoel. The resulting
multicellular structure (embryo) formed as a result of cleavage is called the blastula.

The various planes of cleavage arise according to a fairly definite pattern and in a particular relationship
to each other. Cells typically tend to divide into equal parts (daughter cells). Each new plane of division
tends to intersect the previous plane at right angles. This helps to maintain the spheroidal shape of the
blastomeres. The first cleavage division is often along the polar (animal-vegetal pole) axis of the egg
called the first meridional cleavage. The second division is also vertical and takes place at right angle to
the first cleavage. Thus, both the first and second mitotic divisions are meridional and at right angles to
each other. The plane of the third division is horizontal and either at/ slightly above the equator. ???Further
planes of cleavage
 Cleavage is influenced by the quantity of yolk and also the pattern of distribution of yolk in the egg. This
is because yolk is an inert material and it hinders the progress of cleavage furrows. The division takes
place faster at the animal hemisphere and is relatively slow at the vegetal hemisphere. In fact, if the yolk
is in large amount then the divisions (cleavage furrows) may be incomplete. Thus, cleavage occurs more
readily in the active cytoplasm (yolk-free) rather than the yolk-laden cytoplasm (deutroplasm) of the egg
and is therefore classified into Holoblastic and Meroblastic cleavage types based on whether the cleavage
is total or partial. In placental mammals (including humans) where nourishment is provided by the
mother's body, the eggs have a very small amount of yolk and undergo holoblastic cleavage. Other species,
such as birds, with a lot of yolk in the egg to nourish the embryo during development, undergo meroblastic
cleavage.

1.3.1. Holoblastic cleavage

Holoblastic cleavage means complete division and blastomeres stand distinctly separate from each other.
It is possible only when egg contains small or moderate amount of yolk deposit e.g. microlecithal
(isolecithal) egg and mesolecithal eggs respectively. The holoblastic cleavage can be defined into two
major types depending upon the size of blastomeres viz. equal holoblastic cleavage and unequal
holoblastic cleavage. The equal holoblastic cleavage gives rise to equal sized daughter cells. Whereas in
unequal holoblastic cleavage the blastomeres are of unequal size, the ones which develop in animal
hemisphere are smaller compared to those in vegetal pole. In vegetal hemisphere the blastomeres formed
are of bigger size as they develop slowly due to deposition of yolk. The holoblastic cleavage is further
categorized into four major types based on the symmetry viz. bilateral holoblastic, radial holoblastic,
spiral holoblastic, cleavage and rotational holoblastic.

When the cleavage furrow divides the egg or blastomeres completely, it is called holoblastic cleavage.
The entire egg divides completely into many blastomeres containing yolk. Holoblastic cleavage is
further classified into equal and unequal.

(i) Equal Holoblastic Cleavage – In microlecithal and isolecithal eggs (with evenly distributed yolk)
the entire egg divides producing cells of roughly the same size (or of equal or approx. equal size).
Both the first and second mitotic divisions are meridional and at right angles to each other. The plane
of the third division is horizontal and at the equator. As a result, eight equal blastomeres are formed
which divide further to produce approx. equal-sized blastomeres. E.g. Amphioxus, marsupial and
Placental mammals.

(ii) Unequal Holoblastic Cleavage – In mesolecithal and telolecithal eggs, holoblastic cleavage
produces unequal sized blastomeres, which include many small-sized blastomeres called
Micromeres and a few large-sized yolk-laden blastomeres called Macromeres. For example, in the
eggs of bony fish and amphibians, a moderate amount of yolk is concentrated towards the vegetal
pole. It retards the cleavage furrows in the vegetal pole. Both the first and second cleavage divisions
are meridional and at right angles to each other. The plane of the third cleavage is horizontal but
above the equator. As a result, the blastomeres produced are unequal. The third cleavage, thus gives
 rise to four small and four large blastomeres. The large-sized blastomeres contain more yolk, while
small-sized blastomeres contain less or no yolk. The large blastomeres are called macromeres or
megameres and the smaller ones are called micromeres.

Cleavage based on symmetry: Dependent on the cleavage furrow formation and symmetry the
holoblastic cleavage is further classified as bilateral, radial, spiral and rotational types.

a. Bilateral cleavage

The bilateral cleavage is the result of unequal holoblastic type of division of egg. First bilateral holoblastic
cleavage bisects zygote longitudinally into left and right halves. The blastomeres of successive divisions
are arranged along a single plane, occurring during first meridional cleavage, divisible into two halve
which are mirror images of one another i.e. the blastomeres are arrange in bilaterally symmetrical manner
e.g. vertebrates, some mollusks and some echinoderms.

b. Radial cleavage

The radial cleavage occurs in equal holoblastic division of an egg. In radial cleavage the cell divisions are
equal and symmetrical and occur at right angle to the polar axis of the egg. The blastula is divisible into
two mirror images along axis drawn on any radius i.e. radially symmetrical. It is characteristic feature of
deuterostomes† e.g. echinoderms and Amphioxus.

c. Spiral cleavage

In spiral type of cleavage the resultant cells are placed alternate to each other or in a spiral fashion. This
type of cleavage usually occurs in equal holoblastic type of cleavage, although some undergo unequal
holoblastic cleavage. In this division first two meridional cleavages at right angle to each other giving
rise to four daughter cells as A, B, C, and D, each representing one quadrant of an embryo. These two
cleavages are parallel to the animal-vegetal axis of the zygote. But at 4-celled stage A and C macromeres
meet at animal pole creating animal cross furrow whilst the B and D macromeres meet at vegetal pole
creating vegetal cross furrow (Fig…..). It is found in flatworms, annelids and mollusks known as
protostomes‡. In successive cleavages each division occurs at an oblique angle (not right angle to the
animal-vegetal axis) to give rise to smaller micromeres know as quartet of the mother cell.

†
Animals in which the blastopore develops into anus and mouth is developed on its opposite side.
‡
Animals develop mouth from blastopore.
 Fig. 1.3.1.: Types of holoblastic cleavages

d. Rotational cleavage

In a rotational cleavage normally the first division is along meridional axis giving rise to two daughter
cells. But during second cleavage one of the daughter cell divides meridionally whilst the other
equatorially. The rotational cleavage is found in isolecithal type of mammalian eggs.

Fig. 1.3.1d: Rotational cleavage

1.3.2. Meroblastic cleavage

Meroblastic cleavage means incomplete divisions due the presence of the large amount of yolk preset in
macrolecithal egg (telolecithal and centrolecithal eggs). These are of three main types bilateral meroblastic
cleavage, discoidal and superficial cleavages. The cleavage occurs only in cytoplasm and not in yolk.

a. Discoidal cleavage

Many animals have discoidal eggs which contain very little quantity of cytoplasm as compared to huge
deposition of yolk. In these eggs the cleavage is restricted to only in cytoplasmic disc, hence the name.
The cytoplasmic disc is separated from the yolk by a space called as subgerminal cavity. The discoidal
cleavage occurs in fish, reptiles and bird.

b. Superficial cleavage
Superficial cleavage is limited to a thin surface area of cytoplasm that covers the entire egg. The centre of
the egg is filled with yolk which fails to cleave. Thus, the cleavage does not take place in cytoplasm but
only nucleus is divided through mitosis, known as karyokinesis. These nuclei are referred as energids.
After several rounds of karyokinesis the naked nuclei migrate to the periphery of the egg. At this stage it
is called the syncytial blastoderm because all the nuclei share the same cytoplasm. Cellularzation occurs
at about the 14th nuclear division to create the cellular blastoderm. After this time cells divide
asynchronously e.g. centrolecithal eggs such as insects.

Fig. 1.3.2.: Types of meroblastic cleavages: a. discoidal; b. superficial

1.4. Types of Blastulae

The cleavage advances to the multicellular embryonic stage called blastula. The cells of blastula are
known as blastomeres. At a later phase the blastula develops a central cavity known as blastocoel or
segmentation cavity, surrounded by multicellular layer called as blastoderm. Mainly three kinds of
blastulae are developed in animals e.g. coeloblastula, discoblastula and blastocyst.

Coeloblastula

Coeloblastula is common in eggs having low yolk content. These are spherical and have centric or
eccentric blastocoel. The blastoderm may be made of unilayered or multilayered blastomeres. These are
found in Amphioxus, frogs etc. In some lower invertebrates the coeloblastula itself plays a role of larva
with little modification e.g. in sponges the stereoblastula, parenchymula and amphiblastula which
directly develop into an adult.
 Fig. 1.4a: Coeloblastula and its secondary forms found in lower animals

Discoblastula

Due to the heavy yolk content the cleavage is arrested at vegetal pole and only occurs at animal pole thus
giving rise to polar blastomeres. theses form a flat, topical multilayered disc made of blastomeres called
blastodisc. Blastodisc holds a space just above the yolk mass called subgerminal cavity or primary
blastocoel. The blastodisc remains as a superficial layer detached at the center. The centre of the blastodisc
therefore remains transparent known as area pellucida as compared to the periphery called as area opaca.
The extreme margin of the blastodisc remains attached to the lower yolk mass known as zone of junction.
The blastula formed in this manner is known as discoblastula. It is found in reptiles, birds and monotremes
whose eggs are of macrolecithal type.
 Fig. 1.4b: Discoblastula of birds

Blastocyst

In marsupial and placental mammals the eggs have almost no yolk. The first cleavage divides egg into
two blastomeres out of which one represents animal pole and the other as vegetal pole. Due to the lack
of yolk, the cleavages at vegetal pole are faster as compared to that of animal pole. The cells at the vegetal
pole start growing and dividing vigorously which constitutes the thin outer wall called as trophoblasts
(troph = nourish). Finally at the 32 to 64 celled stages the trophoblasts develop an envelope, enclosing an
eccentric space, where the yolk§ would have been, to develop an eccentric slit like cavity, a blastocoel.
The cells at animal pole are confined to one end (embryonic pole) often referred to as inner cell mass.
The inner cell mass is covered by a layer of Rauber’s cells. The blastula thus developed in form of a cyst
is known as blastocyst. The blastocyst now descends down to uterus and escapes from zona pellucida
and gets implanted on uterine wall. Trophoblasts now start obtaining nourishment from maternal tissue.

Fig. 1.4c: Blastocyst of mammals

1.5. Gastrulation

Gastrulation is a phenomenon of making and shaping the embryo by the course of massive proliferation,
movements and rearrangements of the cells occurred in a simple multicellular ball, the blastula. The cell
movements locate them to their definitive position in the embryo which leads to the formation of basic
germ layers (embryonic layers) which play important role in organogenesis**. These germ layers are
formed due to various cellular activities such as cell motility, their shaping and adhesion known as
morphogenetic movements. During these movements, a new cavity is developed called archenteron
(gastrocoel) which opens to exterior by a blastopore. Invertebrates like poriferans and coelenterates
exhibit very simple gastrulation developing only two germ layers viz. ectoderm and endoderm. These are
referred as diploblastic animals.

§
In mesolecithal and macrolecithal types of eggs the yolk sac usually encloses yolk mass which is utilized in the entire
development of an embryo. However, despite of lack of yolk the cells behave in similar manner indicating evolutionary trend
showing genetic relationship with reptiles.
**
The basic germ layers give rise to different organs of the body referred as organogenesis e.g. neurulation is of
forming neural tissue from ectoderm.
 Fig. 1.5a: Scheme of germ layers in diploblastic and triploblastic animals

Formation of planula larva in coelenterates is classical example of diploblastic gastrula developing into
an adult. In higher animals, three germ layers viz. outermost ectoderm, middle mesoderm and innermost
endoderm, are formed hence they are known as triploblastic. Endoderm forms inner lining of the gut
and many internal organs like kidney, urinary bladder etc. Ectoderm forms external structures like skin,
nervous system, exoskeleton etc. Mesoderm develops into muscles, skeletal and circulatory systems.
Archenteron develops into gut of the animal. In lower invertebrates in which the blastopore develops into
mouth are referred to as protostomes (Phylum: Porifera to Phylum: Mollusca) while in higher animals it
forms anus and mouth develops opposite to it are called as deuterostomes (Phylum: Echinodermata and
above).

Fig. 1.5b: Plannula larva of Sponge

1.5.1. Morphogenetic Movements:

Morphogenetic movements are of two major types viz. epiboly and emboly. Epiboly is a process of
thinning and extension of the epithelium which occurs through different movements of the embryonic
cells as a whole e.g. extension, intercalation, convergence, divergence, etc. whereas, emboly involves the
movements of cells into the interior of embryo e.g. invagination, ingression, infiltration, delamination,
proliferation and involution.
 Extension

Epiboly Intercalation

Convergence/
divergence

Cell Movements in Invagination

Gastrulation
Ingression

Infiltration
Emboly
Involution

Delamination

Proliferation

Fig. 1.5.1: Types of cell movements in gastrulation

Types of movements in epiboly:

Extension: Here the cells get stretched to increase the length of the germ layer.

Intercalations: It is merging of two epithelia in which cells from two adjacent epithelia move between
one another and merge to form a single layer.

Convergence/divergence: It means the intercalation of cells from multilayered epithelium in a highly

directional and organized manner. When cells congregate at one point the process is called convergence
and in case they move away from each other is known as divergence.

Mostly all these movements occur in combinations (Fig. 1.5.1, a).

Types of movements in emboly

Invagination: The layer of epithelial cells migrates inwards and forms a depression. The region from
where the cells invaginate is called the blastopore. It has a dorsal lip and a ventral lip. The dorsal lip is
made of apical cells and the ventral of the basal cells. (Fig. 1.5.1, b). This is seen in Amphioxus, frog.
Ingression: The epithelial cells get separated and migrate inside to form free flowing mesenchymal cells
to get rearranged in the gastrulation process. (Fig. 1.5.1, c). It is seen in echinoderms.

Involution: The epithelial layer rolls down and folds inwards to form a new layer underneath the outer
layer (Fig. 1.5.1, d). It is seen in amphibians and birds.

Infiltration: The cells detach and fall inside the lumen of the embryo (blastocoel) to form loose cell mass
called as mesenchyme (Fig. 1.5.1, e). The mesenchymal cells then spread out quickly to to form a
continuous layer called hypoblast. The blastocoel is now called as archenteron. It is seen in chick and
human.

Delamination: The cells get divided equatorially to form a new layer which ultimately gets separated by
splitting up from the mother layer (Fig. 1.5.1, f). It is seen in mammals.

Fig. 1.5.1: Showing the processes involved in gastrulation of animal embryos

a. Epiboly showing extension, intercalation, convergence and extension

b. Invagination c. Ingression d. involution

e. Infilteration f. Delamination
Gastrulation in protochordates having microlecithal egg (example: Amphioxus)

The gastrulation in amphioxus occurs by the process of invagination forming initially a double layered
cup. The blastocoel reduces by the course of development of the gastrula and the cup shaped formation
holds a new cavity known as archenteron. As the gastrulation advances the opening of the cup constricts
to form blastopore. At this stage the gastrula exhibits outer ectoderm and the inner endoderm with
rudiments of mesoderm (Fig. 151, h).

Fig. 1.5.1, h: Gastrulation in Amphioxus

Gastrulation in amphibian with mesolecithal egg of frog

Due to the presence of adequate yolk the process of invagination of the blastula is restricted only towards
animal pole. Initially an inverted crescentic depression (slit-like) occurs at the equator. The upper margin
of this slit is named as dorsal lip of blastopore. The micromeres at animal pole divide rapidly and
migrate towards the vegetal pole to cover the macromeres through epiboly. During this process the
macromeres shift inside by process of involution and invagination (emboly). The entire mass of
macromeres move inside except a small portion which is visible from the blastopore called as yolk plug.
The embryo becomes double walled gastrula enclosing archenteron opening to exterior with a blastopore.
The macromeres constitute the endoderm lining internal layer of the gastrula and the outer micromere
layer becomes ectoderm. Some cells at the blastopore undergo ingression to form third germ layer called
mesoderm.
 Fig. 1.5.1, i: Gastrulation in frog

Gastrulation in bird with macrolecithal egg.

In discoblastula of birds, the blastoderm remains as superficial layer, the epiblast. The lower layer, the
hypoblast is formed through the phenomenon of several processes†† (as explained by number of theories)
of epiblast cells underneath. The hypoblast divides the subgerminal cavity into dorsal true blastocoel and
ventral, the archenteron. It has been stated that the cells forming endoderm migrate from posterior to
anterior direction of the embryo. The formation of mesoderm initiates with the origin of primitive streak
developed at posterior end along the mid-dorsal line of the discoblastula. The primitive streak later grows
forward and elongates. The thickening of the primitive streak occurs due to convergence of epiblast cells
due to the presence of underlying hypoblast cells. There is a furrow lying mid-dorsally to the primitive
streak called primitive groove which forms primitive folds at the lateral edges. At anterior end of the
primitive groove is a compact mass of the cells called Hensen’s node having a central depression, the
primitive pit. At posterior end, the primitive streak has a plate called primitive plate. As the primitive
streak elongates towards the anterior end, the area pellucida gets thickened and forms elliptical embryonic
shield. The mesenchyme cells proliferate rapidly near the primitive groove and migrate to occupy
blastocoel present between epiblast and hypoblast. They continue migrating antero-laterally unto the limit
of primitive streak. These mesenchyme cells constitute the mesoderm. With the advancement of the
embryo, the primitive streak disintegrates and recedes backwards dragging Hensen’s node along. The
Hensen’s node goes on releasing behind the prechordal and notochordal cells (modified epiblast cells)
which ultimately develop into nerve cord and notochord respectively. The primitive streak disintegrates
in such a manner that its remnants restrict merely to the tail and cloacal region of the hatchling.
Nevertheless, the epiblast and hypoblast later become ectoderm and endoderm, respectively. The

††
It was explained by many experts as theory of ingression (Peter, 1983); poly-ingression (Pasteels, 1945); delamination
(Spratt, 1945); invagination (Jacobson, 1938) or involution (Patterson, 1909).
mesoderm free area lying anterior to primitive streak is called as proamnion. The proamnion region plays
a role in the formation of a distinct head known as cephalization (Fig. 1.5.1, j).

Fig. 1.5.1, j:

Gastrulation in mammal with alecithal egg

It has been believed that the trophoblasts after hatching out of zona pellucida absorb the surrounding
uterine fluid for their nourishment to grow and multiply faster. Trophoblasts play important role in
formation of a placenta. In mammals, the gastrulation initiates with the horizontal division of inner cell
mass of blastocyst by the process of delamination to give rise to upper epiblast and the lower hypoblast.
The hypoblast grows downwards to across the inner lining of trophoblasts to form endoderm. Endoderm,
eventually meets mid-ventrally to enclose blastocoel to form archenteron or yolk sac. The Rauber’s cells
now disintegrate to expose epiblast which develops into polar disc or embryonic disc by multiplication
and rearrangement of the remaining inner cell mass. The upper cell-layer of this embryonic disc then
merges with trophoblasts to form ectoderm which is then called trophectoderm. Trophectoderm also in
due course of time extends over the whole embryo to form outer germ layer.

Development of primitive streak and subsequent formation of mesoderm initiate after the formation of
trophectoderm. Formation of primitive streak involves rapid multiplication of the cells in the region
causing thickening of the area. Initially, an elongated primitive streak originates at the caudal end in the
form of a crescentic outgrowth and extends towards the cephalic end of the embryo. Due to the crowding
of cells along the sides of the primitive streak, there occurs ingression of some cells to form mesenchyme
cell layer between the endoderm and trophectoderm. The mesenchyme cells later start multiplying and
migrating antero-laterally to occupy the space between ectoderm and endoderm to modify into a middle
germ layer, the embryonic mesoderm. The ingression of mesenchyme cells also occurs outside the
embryonic disc to form extraembryonic mesoderm. The extra-embryonic mesoderm plays an important
role in the development of extraembryonic (foetal) membranes (Fig. 1.5.1, k).
 (i) Blastocyst with basic inner cell mass and without (ii) Blastocyst showing commencement of
endodermal layer endoderm formation from hypoblast growth within
the trophoblast to form yolk sac

(iii) Endodermal layer extends throughout the entire

trophoblastic layer internally and meet at the bottom
enclosing the imaginary yolk sac. The development of
hollow yolk sac gives the evidence of origin of mammals
from the ancestral reptiles in which the egg had ample
yolk and yolk sac developed as a rule. The epiblast and
hypoblast form compact blastodisc.
(iv) The primary yolk sac is formed without yolk
and the space is empty inside. Meanwhile
extraembryonic mesoderm originats from embryonic
mesoderm that advances between the endoderm and
ectoderm.

Fig. 1.5.1, k: Gastrulation in blastocyst of mammals (i, ii, iii & iv)

1.6. Coelom – Formation and Types

Coelom is a fluid filled body cavity formed in triploblastic animals which is internally lined by mesoderm.
Coelom protects the internal organs acting as a cushion against accidental shocks. It also allows organs to
grow and move internally. In lower animals like earthworm the coelom along with the body-fluid works
as a skeletal system.

1.6.1. Formation of Coelom (Schizocoely and enterocoely)

Coelom is developed from mesodermal layer. In lower triploblastic animals like protostomes the
mesoderm splits into inner and outer layers to form the body cavity, the phenomenon is known as
schizocoely. The inner layer restricted to endoderm is called as splanchnic mesoderm and the outer which
confines to ectoderm is called as somatic mesoderm. The union of splanchnic mesoderm with endoderm
is known as splanchnopleure while the somatic mesoderm with ectoderm is called somatopleure.

In deuterostomes the mesoderm out-pockets from the gut endoderm and later gets plugged to form hollow
structure which later expands to occupy the inner space forming a coelom. This phenomenon is called as
enterocoely. However, the mesoderm in enterocoely develops splanchnic and somatic layers and
eventually get associated with ectoderm and endoderm to form splanchnopleure and somatopleure
respectively.
 Fig. 1.6.1a: Schematic diagram of coelom formation

1.6.2. Types of Coelom

The coelom which is enclosed entirely in mesodermal epithelium is called as true coelom and the animals
having true coelom are referred as eucoelomates.

Fig. 1.6.1b. Types of coelom in (a) Acoelomates (b) Pseudocoelomates and (c) Eucoelomates

Nevertheless, not all triploblastic animals develop the coelom in these manners. Those which do not
develop coelom at all are referred as acoelomates e.g. platyhelminths (flat worms; Planaria, Taenia). In
round worms the coelom is developed by shifting of mesoderm towards ectoderm hence holding cavity
between endoderm on inner side and mesoderm on the outer. The coelom thus formed is called as false
coelom and the animals having such coelom are known as pseudocoelomates e.g. Aschelminthes (round
worms; Ascaris).

1.7. Extraembryonic Membrane

Extraembryonic membranes are developed from the germ layers of embryo itself but not incorporated
within the embryo. They develop outside the embryo and help in protection from desiccation, infection
and shock. They supply of nutrients, exchange of gases like oxygen and carbon dioxide and remove waste
matter. Hence they are known as extraembryonic membranes. Principally, the extraembryonic membranes
are of four type viz. chorion, amnion, yolk sac and allantois. The vertebrates which develop amnion are
called as amniotes (reptiles, birds and mammals) and others are called as anamniotes (fishes and
amphibians). In anamniotes, several primitive types of accessory embryonic membranes are developed in
addition to yolk sac. e.g. in some fishes a pseudochorion and a pseudoamnion (poeciliid fishes) is
developed from pericardium in neck region of embryo.

Fig. 1.7: Schematic diagram showing the layout of extraembryonic membranes found in amniotes

1.7.1. Yolk sac

In anamniotes and amniotes, the yolk is enclosed in a yolk sac which develops outside the embryonic
tissue as an extraembryonic membrane. The yolk sac develops from the endodermal layer, continuous
with that of embryonic gut, beneath the vitelline membrane enclosing the entire yolk mass. In some
vertebrates, like teleosts and amphibians, the yolk is incorporated with the embryonic tissue present in
form of a little bulge at the abdominal region and hence is not an extraembryonic membrane. It is called
endogenous yolk sac‡‡. Whereas in polylecithal eggs in sharks, reptiles, birds and egg-laying mammals,
due to the presence of large amount of yolk the cleavage at vegetal pole is restricted. Hence the yolk sac
is developed from external growth of endoderm which extends and encloses the yolk mass to form yolk
sac outside the embryo. The phenomenon in which yolk is digested and absorbed through the
extraembryonic membranes is known as exogenous yolk sac. Usually it is developed in eggs having large
amount of yolk e.g. oviparous sharks, reptiles, birds and mammals. But it is also found in alecithal eggs
in case of placental mammals and play different role. The yolk is digested by endodermal epithelium and
is absorbed by network of capillaries around yolk sac which collect blood in vitelline veins and is
ultimately supplied to embryo. As the yolk is utilized the yolk sac diminishes in size and eventually is

‡‡
In teleosts (bony fishes) and amphibians, yolk is divided with cleavage forming macromeres. However, these macromeres
laden with yolk get concentrated in abdominal region bulging out ventrally as a yolk reserve termed as endogenous food supply.
retrieved inside the abdomen at mid gut area. Its remnant is found as a temporary scar or diverticulum
which in man is known as Meckel’s diverticulum.

Fig. 1.7.1a: Endogenous yolk sac in a teleost fry & frog

Oviparous anamniotes lay mesolecithal eggs having ample food reserve so as to carry out the embryonic
development outside the parental body, in open aquatic medium. Water medium avails exchange of gases
and release of excreta except nutrition. Therefore in oviparous anamniotes, only yolk sac is formed as an
extraembryonic membrane. The primitive yolk sac is made of endoderm, mesoderm and ectoderm held
compact without intervening space. It is found in some cyclostomes, most elasmobranchs (cartilaginous
fish) and teleosts.

Fig. 1.7.1b: Cross sections through exogenous yolk sac (a) primitive (shark) and (b) advance type (birds)

In dogfish, with the development of embryo the yolk is isolated in yolk sac which is connected to embryo
by a yolk stalk containing the vitelline veins and arteries. The yolk is utilized by new born pups of dogfish
until they are able to capture food, their own. In viviparous anamniotes, like dogfish the yolk sac also
helps in absorbing uterine milk secretion after yolk is utilized by forming a sort of loose placenta against
the uterine wall of the mother.

(a) (b) (c)

Fig. 1.7.1c: Exogenous yolk sac in (a) shark, (b) chick and (c) placental mammals

Amniotes, except placental mammals, develop advanced type of yolk sac around huge yolk reserve in
their eggs. The splanchnopleure grows slowly around the yolk mass to enclose it entirely forming a yolk
sac. The inner lining of yolk sac develops numerous fingerlike outgrowths into yolk mass called as yolk
sac septa and the outer surface is innervated by numerous vitelline veins and arteries making it highly
vascularized. The splanchnopleure constrict at the midgut region to form a narrow yolk duct constituting
the yolk stalk.
In placental mammals, the yolk sac is developed in a similar manner as in reptiles and birds despite the
eggs being yolkless. The trophoblasts of the blastocyst, enclosing the empty space as a yolk sac, are lined
internally by extraembryonic endodermal layer. It becomes highly vascularized with vitelline blood
vessels. Yolk sac in placental mammals play important role in the formation of the part of umbilical cord
and highly vascularized surface lying close to maternal tissue forming a placenta. The yolk sac plays
important role in transport of uterine milk to embryo and also helps in its respiration.

In ovo-viviparous reptiles and marsupials, yolk sac associates with the chorion to form chorio-vitelline
placenta. In other mammals like Lagomorpha, Insectivore, Chiorptera and Rodentia it plays a temporary
role during gestation.

1.7.2. Formation of amnion and chorion:

Both the amnion and chorion are derived from an extraembryonic membrane known as somatopleure.
Number of double walled amniotic folds of somatopleure originated from the base of embryo which is
named on the basis of their corresponding positions as lateral folds, head fold and tail fold. The head fold
appears first followed by two lateral folds which grow upwards and backwards. Finally the tail fold
develops and grows upwards as well as anteriorly to meet the prior three folds. Eventually all the folds
meet each other and fuse at a slightly posterior point. This point before closure is referred to as amniotic
umbilicus. After the fusion of the folds, the outer and inner layers are separated as chorion and amnion
respectively.

1.7.3. Amnion

Amnion, the primary extraembryonic membrane of amniotes, is a protective membrane that surrounds the
embryo forming a sac of fluid. It is an impermeable membrane retaining the fluid for the free floating of
embryo for its growth and development. Amniotes being terrestrial organisms, amnion resembles the
primordial aquatic environment of anamniotes from which the former have evolved.

The amnion is formed of two layers, the outer somatic mesoderm and the inner ectoderm. The cavity is
known as amniotic cavity and is filled with amniotic fluid. Amniotic fluid being viscous, it not only
provides free floatation of embryo for its even growth but also it functions as an efficient shock-absorber
to protect it from mechanical injury. The free floatation of embryo ensures to avoid contact with the lateral
walls or shell preventing unwanted deformity. Also it protects from the desiccation of embryo. The amnion
is connected to the embryo at ventral hollow stalk called as somatic umbilicus.
 Fig. 1.7.3: Formation of amnion and chorion: (a) formation of amniotic folds; (b) separation of chorion from amnion

1.7.4. Chorion

The outermost layer of extraembryonic membranes that is principally involved in respiration in birds and
has many other functions in mammals. During the formation of amnion the amniotic folds meet at amniotic
umbilicus and the two layers are separated as inner amnion and the outer chorion. This is also referred as
false amnion. The wall of the chorion is also made of two layers, the outer being ectoderm and the inner,
somatic mesoderm.

1.7.5. Allantois

Allantois is the outgrowth of the ventral part of the hind gut of embryo. Proximally it permanently remains
as embryonic part but distally it grows towards exterior and unites with splanchnic mesoderm to form
splanchnopleure. In viviparous amniotes, the allantois expands further to lie close to outer porous egg-
shell and helps in respiration and deposition of wastes from gut and kidney until the egg hatches. However,
the Allantois does not come in direct contact with shell due to middle layer of chorion. In viviparous
amniotes, the origin and development of allantois is similar but it forms complex with chorion to impart
formation of chorio-allantoic placenta. The allantois becomes highly vascularized and helps in nutrition
as it draws nourishment provided by the maternal tissues. The allantois in amniotes, except birds, becomes
obliterated and is pinched off during the completion of gestation and its proximal part gets retrieved into
abdominal cavity to form into urinary bladder.

Fig. 1.7.5: Allantois incorporated with chorion to enhance the gaseous exchange of embryo
1.8. Types of Placentae Based on Morphology, Implantation and Histology

The placenta is a characteristic of viviparous vertebrates and does not occur in egg laying animals. When
the eggs of vertebrates contain insufficient yolk reserve, the development of embryo is augmented by
peculiar foeto-maternal associations. There are various foeto-maternal associations which accomplish
nourishments of their embryos. The association intensity ranges from a very simple type as in lower
vertebrates to a highly complex form as in mammals known as eutherians. The simple association is found
in goodeid fish in which, trophotaeniae (anal structure) is developed and in anablepiid fish the maternal
villi are in contact with fetal belly sac. Some fishes exhibit modified fin folds helping to nourish embryo,
establishing association with maternal tissue. The larvae of frog which develop in pouches on the back
are attached to the parent with extensions of their gills for their nourishment. Similarly, gill filaments of
larvae of viviparous salamander, Proteus establish the contact with uterine wall. Some reptiles (e.g. garter
snakes) are viviparous and develop a rudimentary placenta.

Placenta in mammals:

The egg laying mammals (monotremes) have ability to lactate but have no placenta. They lay eggs with
leathery shells, which the females then incubate in a pouch. The mammals other than egg laying mammals
are known as therians (marsupials and eutherians) which develop various types of placentae depending
upon their morphological and histological characteristics and intimacy of foeto-maternal association.
Marsupials, referred as metatherians, are the pouch bearing mammals in which the underdeveloped
embryos are reared until they are completely developed. Eutherians are viviparous mammals and form
true placenta as a foeto-maternal association and hence commonly referred as true placental mammals. In
mammals the trophoblasts are sycytialized that coordinate with the biomaolecular interactions between
the fetus and mother. Major hormones are also produced by placental syncytiotrophoblasts.

The mammalian placenta is classified on the basis of whether the wall of allantois comes in contact with
the chorion or not. Basically, the combination of extraembryonic membrane viz. chorion, allantois and
yolk (vitelline) sac plays important role in development of placenta from foetal side. These
extraembryonic membranes combine with the maternal reproductive tract to form a close association of
the tissue enabling them to exchange life essential materials for the growing embryo. The vital components
such as oxygen, carbon-dioxide, hormone, nutrition and toxicants are exchanged at placenta through the
network of blood capillaries without the mixing of maternal blood with that of the embryo. There are three
major types of placenta in therians called as chorio-vitelline, chorio-allantoic and chorion alone. These
extraembryonic membranes combine with maternal mucosa to form a highly vascularized and close
opposition of a structure called placenta. Placenta is also defined on the basis of the implantation of
embryo in the uterine wall.

Chorio-vitelline placenta:
The marsupial mammals like kangaroos have a rudimentary and short-lived placenta which, in most
marsupials, structurally and functionally is different from the typical eutherian placenta. In marsupials
allantois remains small and does not fuse with chorion to form placenta. Instead yolk sac develops large
in size and combines with chorion to give rise to chorio-vitelline or yolk-sac placenta. The blood is
supplied through vitelline blood vessels to placenta from embryonic side. Chorion does not form villi to
construct placenta and remains smooth surface in close opposition with maternal vascular mucosa, the
endometrium. Placental nourishment of marsupial young is negligible compared to nourishment from the
milk obtained in the pouch.

Chorio-allantoic placenta

In this case the yolk sac remains small or rudimentary and allantois grows large and combines with chorion
to give rise to chorio-allantoic or allontochorion placenta. It is made up of outermost layer of chorionic
trophoblasts, middle mesoderm of chorion and allantois and the inner layer of allantoic endoderm. The
mesoderm is a thick layer made of fused mesodermal layers of chorion and allantois. This layer contains
numerous spaces filled with embryonic blood. The allantois becomes highly vascular and takes part in
foeto-placental circulation. The chorion develops several villi to establish more close association with
maternal mucosa, for enhancing the exchange of vital matter in placenta. From maternal side the uterine
mucosa and submucosa form the union with the allantochorionic villi to make a placenta. This type of
placenta is developed in some marsupials and all eutherians.

Chorionic placenta

When the association of allantois with chorion is insignificant or absent then structure is referred as
chorionic placenta.

The placenta is defined on the basis of the type of implantation of embryo in the uterine wall. It depends
upon the position and mode of attachment of embryo. There are three modes of implantation of embryo
in the uterine wall viz. central, eccentric and interstitial.

i. Central or superficial implantation:

The embryo remains in the centre of the uterine lumen and extraembryonic membranes make superficial
contact with uterine mucosa. This type of implantation is common amongst the lower mammals e.g.
marsupials (Perameles and Dasyurus) and eutherians (sheep, dog, pig, rabbit, cow, cat, etc.)

ii. Eccentric implantation:

The embryo lies on one side of the endometrial wall and is embedded by the formation of epithelial folds.
In other words the uterine wall forms the cavity on the endometrial wall in which the embryo lie
embedded. Since the embryo is partially buried inside the uterine wall it lies on one side of the uterine
lumen thus it is called as eccentric implantation. The embryo, however, is not covered entirely by the
uterine epithelium. This type of implantation is found in mouse and rat.

iii. Interstitial implantation:

The embryo is completely buried inside the uterine mucosa below the epithelium. Therefore it is
completely covered by endometrial tissue. This type of implantation is found in human, chimpanzee and
guinea pig.

The placentae are further classified on three principle bases viz. (i) morphological characteristics, (ii) type
of implantation of embryo in the uterine wall and (iii) intimacy of foeto-maternal tissue in placenta and
(iv) histological structure of the placenta.

Classification based on morphological characteristics of placenta:

The types of placentae are defined on the pattern of the villi formation on the chorion of the blastocyst.
They are of five main types.

i. Diffused placenta

The villi are distributed uniformly on the surface of the chorion except the terminal ends. This is found in
mare, pig, etc.

ii. Cotyledonary placenta

The villi are developed in patches on the chorionic surface whilst the remaining part of the chorion is
smooth. These patches are called cotyledons. This type is found in sheep, goat and cattle.

iii. Intermediate placenta

This is the combination of diffused and cotyledonary placenta. There are some isolated villi present
between the cotyledonary patches. This type of placenta occurs in camel and giraffe.

iv. Zonary placenta

The villi are developed along a equatorial belt on the blastocyst. This belt is usually elliptical in shape.
This type is seen in carnivores like dog, fox, mongoose, tiger, lion, cat etc.

v. Discoidal placenta
The villi are confined in a disc like area, eccentrically, on the blastocyst. It is found in insectivores rat,
mouse, bat, rabbit and bare. In monkeys, two discs are developed on the blastocyst, and therefore referred
as bidiscoidal placenta e.g. human, mouse etc.

Classification of placenta based on intimacy of foeto-maternal tissue:

Intimacy of the union of foeto-maternal tissue defines three kinds of placenta, viz. non-deciduate,
deciduous and contra-deciduous.

i. Non-deciduous/deciduate placenta

In the mammals, in which the superficial placentation occurs the villi are just pushed into the depressions
of uterine mucosa. In this association the chorionic surface lie close but and endometrium do not fuse with
each other. The nutrients, gases and waste products are passed through all six placental layers of tissue.
During parturition, the chorionic villi are just drawn out from the endometrial depressions without causing
any damage to the uterine wall, hence no bleeding occurs. Found in cattle, pigs etc.

ii. Deciduous/deciduate placenta

In this type, the foeto-embryonic association more close and compact as compared to non-deciduate
placenta. The chorionic trophoblasts penetrate deep inside the eroded uterine mucosa. As a result the
uterine damage and excessive bleeding occurs at the time of parturition. The maternal tissue which is
expelled at the time of parturition is known as duciduae. The more intimate placental association enhances
the exchange of nutrition and gases from maternal tissue. The deciduous placenta is also referred as to
true placenta. It is seen in eutherians.

iii. Contra-deciduous placenta

In this type of placenta the maternal as well as the foetal tissue is lost during the parturition. The tissue is
absorbed by the maternal leucocytes in situ instead of expelling out. It is observed in Parameles (desert
bandicoot) and Talpa (mole).

Classification of placenta based on histological characteristics:

Normally the placental wall, histologically, is composed of six layers of tissue viz. these are known as
barriers between the maternal and foetal blood streams. With the improvement in the placenta the number
of barriers tends to reduce. With the reduction of the number of barriers the foeto-maternal association
becomes more and more intimate. This is known as Grossor’s classification of placenta.

a. Endothelial wall of maternal blood vessels,

b. Connective tissue around maternal blood vessels.
c. Uterine epithelium
d. Chorionic epithelium
e. Chorionic connective tissue
f. Blood vessels in the wall of chorion

Grossor’s classification of placentae e.g. 1) Horse 2) Sheep; 3) dog and 4) human

This thickness of the placenta may be decreased by removal of some of the layers. The histological type
of placenta is defined on the basis of the layer which is missing from it.

1. Epithelio-chorial placenta:

In this case, the chorial epithelium and uterine epithelium unite to form a placenta known as epithelio-
chorial placenta. The chorionic villi are pushed into the depressions of the uterine wall. This type of
placenta is non-deciduous and retains all six layers of placenta. Therefore this placenta is considered as
primitive type from which others have been derived. It is seen in ungulates and marsupials.
2. Synepithelio-chorial placenta

The uterine epithelium is lost in the process of development of this type of placenta. The chorion becomes
directly in contact with the endometrial connective tissue (desmos) instead of uterine epithelium.
Therefore only five layers are retained in this placenta. Due to more intimate association the damage in
the uterine wall occurs during the parturition. It is found in buffalo, cow, sheep, goat, camel, and giraffe.

3. Endothelio-chorial placenta

In this type of placenta, the uterine epithelium and the connective tissue are lost, and it is made of only
four layers. Therefore the chorionic epithelium becomes direct in contact with the endothelial wall of
maternal blood vessels. It is seen in carnivores like cat, tiger, lion, fox, dog, mongoose, walrus etc.

4. Haemo-chorial placenta

In this the endothelial wall of blood vessels also disappears and only three layers of barriers remain. The
chorionic villi are surrounded by maternal blood sinuses. In these sinuses the maternal blood is supplied
through the uterine arteries and collected by the uterine veins. It is found in primates like ape, lemur,
woman; many insectivores like bats and rodents like mouse and rat.

5. Haemo-endothelial placenta

In this case all three maternal barriers and two foetal barriers viz. epithelium and connective tissue of
chorion are lacking. The number of barriers is reduced to just one therefore making the physiological
exchange more efficient. The endothelial wall of chorionic blood vessels bathed directly into blood sinuses
of the uterus. This type of placenta occurs in guinea pig, rabbit and rat.

Learner’s space:

However, the cavity present within the body of sponges is known as spongocoel and that of coelenterates
which performs many functions such as digestion, circulation, respiration, excretion etc. In coelenterates
the archenteron modifies as body cavity called as coelenteron. Coelenteron, in addition to the functions
mentioned in spongocoel holds gonads and involves in reproduction. However, in lower invertebrates
the archenteron which act as body cavity which is not homologous to the coelom of higher animals.

Models

Internet animations
Reference Books

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11. Shumway, W., 1935. Introduction to Vertebrate Embryology. A textbook for College and University. 3rd
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Websites visited:

http://www.biologydiscussion.com/notes/structure-functions-and-types-of-mature-sperm-in-animals-biology/768

http://www.ncbi.nlm.nih.gov/pubmed/6409171

Boundless. “Cleavage, the Blastula Stage, and Gastrulation.” Boundless Biology. Boundless, 08 Aug. 2016.
Retrieved 18 Aug. 2016 from https://www.boundless.com/biology/textbooks/boundless-biology-textbook/animal-
reproduction-and-development-43/fertilization-and-early-embryonic-development-242/cleavage-the-blastula-
stage-and-gastrulation-899-12150/